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Coleoptera, Hydrophilidae) in South-Western Iberian Peninsula

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Coleoptera, Hydrophilidae) in South-Western Iberian Peninsula PATIERNS OF SMALL AND LARGE-SCALE SPATIAL DISTRIBUTION OF COPROPHILOUS SPHAERIDIINAE (COLEOPTERA, HYDROPHILIDAE) IN SOUTH-WESTERN IBERIAN PENINSULA E. ROMERO-ALCARAZ *, F. SÂNCHEZ-PINERO ** & J.M. ÂVILA* RÉSUMÉ Lors d'un cycle annuel, on a utilisé des pièges appâtés par des excréments de bovins, pour étudier la distribution spatiale de quelques espèces de Sphaeridiinae (Col. Hydrophili­ dae) dans deux biotopes (prairie et savane), d'une ferme d'élevage située à Chiclana de la Frontera (Cadiz, Espagne). Toutes les espèces capturées (14 au total) se trouvaient dans les deux biotopes considérés ; on a ainsi observé une similitude très élevée. Bien que le nombre moyen des espèces par piège soit plus grand dans les prairies, la diversité est très semblable entre les biotopes. La distribution des espèces les plus abondantes est analysée à deux niveaux : entre pièges (niveau comportemental) et entre biotopes (niveau écologique). Les espèces se distribuaient au hasard pendant une grande partie de l'année, et quand il y a eu agrégation, celle ci fut faible. Au niveau écologique, seul Sphaeridium scarabaeoides a montré une préférence significative pour les prairies, alors que le reste des espèces se distribuait également dans les deux biotopes. On propose que de tels modèles de distribution traduisent le caractère généraliste des Sphaeridiinae autant à petite qu'à grande échelle. SUMMARY Pitfall traps baited with cattle dung were used during a year-cycle to study the spatial distribution of severa! species of Sphaeridiinae (Col. Hydrophilidae) in two biotopes (grassland and savanna), from a cattle farm in Chiclana de la Frontera (Cadiz, Spain). Fourteen species were collected, ali of them inhabiting both grassland and savanna, thus, similarity was high between biotopes. The mean number of species per trap was higher in grassland, nevertheless diversity was very similar in the two biotopes. The spatial distribution of the most abundant species was analysed at two different scales: among close traps (distribution at behavioural scale), and habitat selection between grassland and savanna biotopes (distribution at ecological scale). The species were randomly distributed at small spatial scale during most part of the year, aggregation being weak when it occurred. At ecological scale only Sphaeridium scarabaeoides showed a significant selection for grass­ land, whereas the other species were equally distributed in both biotopes. We propose that such distribution patterns reflect the ecological generalism of Sphaeridiinae, acting from narrow to wide (geographical) scales. * Departamento de Biologfa Animal y Ecologia. Universidad de Granada. E-18071, Granada, Spain. E-mail: [email protected] ** Dept. Environmental Science & Policy. University of California, Davis. Davis CA, 95616, USA. Rev. Écol. (Terre Vie), vol. 56, 2001. 37 INTRODUCTION One of the most striking facts concerning coprophilous insects communities is their buge diversity and the high abundance of sorne taxa: according to Hanski & Koskela (1979) more than 50 species and 1 000 individuals may coexist in one individual dung patch. This fact is related to the complex nature of the resource "dung" due mainly to two circumstances: its patchy distribution within the habitats, and the differences (chemical composition, temperature, size ...) among individual patches. Although sorne factors, different from those of ecological, geographical, historical or phylogenetical nature, may influence local diversity of coprophagous beetles (Lobo, 1997), it appears clear that segregation due to interspecific competition is relevant in structuring and diversifying dung beetle communities, although it has been poorly experimentally studied (Hanski, 1991). Species usually segregate using "temporal and spatial refuges" (Giller & Doube, 1994 ), along the time axis (different moments of occurrence) (Finné & Desiére, 1971; Hansk:i & Koskela, 1979; Estrada et al. , 1993), along the spatial axis (inhabiting different places or habitats) (Hanski & Koskela, 1979; Lumaret, 1983), or along the fe eding axis (feeding on severa} types or on different states of degradation of resources) (Landin, 1967; Finné & Desiére, 1971; Koskela, 1972; Sânchez-Pifiero & Avila, 1991; Estrada et al. , 1993; Sowig & Wassmer, 1994). Specialization is, consequently, a valuable way to use such a variable resource. Many other factors may be infiuential in the separation of the species, but the spatio-temporal variation in the occurrence of the species may explain up to 72 % of the differences among dung beetle assemblages (Stevenson, 1982). Thereby, diminishing interspecific competition by means of the allotment of resources among the species, allows the coexistence of species, and therefore a high diversity is favoured. As a consequence of segregation, different patterns of spatial distribution can be found among dung beetles, aggregation dynamics being usual for those more specialized species (Hirschberger, 1998; Palestrini et al. , 1998). Two main scales can be defined to understand spatial distribution of dung beetles. The first of them was called behavioural scale by Hanski (1980b), which involves short-distance movements between patches (i.e. different dung pats) within the same habitat. The second is the scale called ecological (Hansk:i, 1980b), that is, involving movements between patches (i.e. differentbiot opes) far away one from each other. Although these two scales are useful from a practical point of view, we cannot assure that two types of movements (short distance and long distance) do really occur, but rather it is possible that migration from one habitat to another is nothing but an accumulation of short-distance m.ovements, and that habitat-selection is not a consequence of the conditions of the biotope as a whole, but simply of environmental conditions at the microhabitat scale (Hanski & Koskela, 1979); moreover, such a selection may refiect just severa} feeding specializations that promote the distribution patterns of populations. Nevertheless, sorne authors (Stevenson, 1982; Lumaret, 1983) consider the macrohabitat to be a major factor contributing to separate populations of different dung species, and from this point of view, it would be possible that migrations at ecological scale do really occur. On the other band, the spatial patterns of the species may not be the same for males and for females, due to the probably different requirements of both sexes (locating appropriate sites for nesting and development of the larvae, in the case of females, or mating competition, for males). Most of the studies above dealt with Scarabaeoidea dung beetles. They constitute the majority of taxa and individuals in dung and are usually specialized - 38 - in the use of severa! types or states of dung. Nevertheless, Sphaeridiinae (Hydrophilidae) are one of the most abundant groups in temperate dung beetle communities (Desiére & Thomé, 1977; Hanski, 1980d; Franch et al. , 1990), adults being coprophagous and larvae predators (Boving & Henriksen, 1938; Mohr, 1943; Sanders & Dobson, 1966; Hansk:i, 1980a, 1980c, 1980d; Sowig et al., 1994). Contrary to Scarabaeoidea, they show a wide trophic generalism (Mohr, 1943; Landin, 1957; Rainio, 1966; Landin, 1967; Sowig & Wassmer, 1994) and wide distribution patterns (Chiesa, 1959; Hansen, 1987, 199 1). However, little is known about the biology and ecology of Sphaeridiinae in Iberian Peninsula (Garcfa­ Camporro & Arradon, 1984; Franch et al., 1990; Romero-Alcaraz et al. , 1997). Taking this into account, in this paper we study the spatial distribution patterns of a non-specialized guild of coprophagous beetles. We expect them not to follow aggregation dynamics due to their trophic generalism, then acting as a large, weil defined, piece of the dung insect community. Thus, the main goals of this paper are: 1) to describe the distribution pattern at small (behavioural) scale of the species of Sphaeridiinae in a location in southern Iberian Peninsula; 2) to test the existence of habitat selection (distribution at ecological scale) by the species of Sphaeridiinae and 3) to explain such distribution patterns according to different ecological fe atures in the study area. STUDY AREA The study area is located in south-western Iberian Peninsula, in a coastal location of Cadiz province. The field work was undertaken at a cattle farm in the town of Chi cl ana de la Frontera. Cattle rnoves free during the day ali over the farm, and therefore the cattle pressure and the density of dung is rather homogeneous ali around the area. Samples were taken from two different biotopes: grassland and sa vanna. The main differences between both biotopes are the nature of the soil and vegetation. In the grassland, soils are mainly clayey (clay constituting more than 35 % ), stones being absent, although calcareous nodes are rather common. The soil is very plastic when wet, but becomes rather hard when dry. This type of soil shows a very poor drainage, so areas in which it occurs are usually floodedduring rainy periods. In this biotope, herbaceous and nitrophilous pastures are widely dominant. On the other band, soil from savanna is not weil developed and, which is more relevant, it is covered with a thick sandy stratum, 1-1.5 meters deep. As in the case of grassland, no stone can be found in this biotope. Due to its sandy nature, soil from savanna bas a very high drainage, and is subjected to quick changes of temperature. With regards to vegetation, savanna is a "dehesa" biotope, where pastures are dominant,
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